Advancing the causal theory of natural selection

The Modern Synthesis embodies a theory of natural selection where selection is to be fundamentally understood in terms of measures of fitness and the covariance of reproductive success and trait or character variables. Whether made explicit or left implicit, the notion that selection requires that some trait variable cause reproductive success has been deemphasized in our modern understanding of exactly what selection amounts to. The dissertation seeks to advance a theory of natural selection that is fundamentally causal. By focusing on the causal nature of natural selection (rather than on fitness or statistical formulae), certain conceptual and methodological problems are seen in a new, clarifying light and avenues toward new, interesting solutions to those problems are illustrated. First, the dissertation offers an update to explicitly causal theories of when exactly a trait counts as an adaptation upon fixation in a population and draws out theoretical and practical implications for evolutionary biology. Second, I examine a case of a novel character that evolves by niche construction and argue that it evolves by selection for it and consider implications for understanding adaptations and drift. The third contribution of the dissertation is an argument for the importance of defining group selection causally and an argument against model pluralism in the levels of selection debate. Fourth, the dissertation makes a methodological contribution. I offer the first steps toward an explicitly causal methodology for inferring the causes of selection—something often required in addition to inferring the causes of reproductive success. The concluding chapter summarizes the work and discusses potential paths for future work.